WOOD ANATOMY of the TRIBE DIPTERYGEAE with COMMENTS on RELATED PAPILIONOID and CAESALPINIOID LEGUMINOSAE Peter Gasson
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IAWA Journal, Vol. 20 (4), 1999: 441-455 WOOD ANATOMY OF THE TRIBE DIPTERYGEAE WITH COMMENTS ON RELATED PAPILIONOID AND CAESALPINIOID LEGUMINOSAE by Peter Gasson Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AB, United Kingdom SUMMARY The wood anatomy of all three genera in the tribe Dipterygeae, Dipteryx, Pterodon and Taralea is described here in a systematic context. Dipteryx and Pterodon have short, narrow, storied rays and fine intervessel pitting, whereas Taralea has narrow non-storied rays and much finer intervessel pitting. The wood of T. casiqui arensis is much more similar to that of Dipteryx and Pterodon than to its congeners, and a new combination based on both wood and morphological features is being published in Kew Bulletin (Lewis & Gasson, in press). The Dipterygeae may form an outlying group to the Dalbergieae, although its nearest affinity has been thought to be with Monopteryx (Sophoreae). Here, the wood anatomy of the three genera is compared with that of Sophoreae, Swartzieae and Dalbergieae. Key words: Dipterygeae, Dipteryx, Pterodon, Taralea, wood de scriptions. INTRODUCTION The tribe Dipterygeae (Leguminosae, Papilionoideae) comprises three genera from Central and South America: Dipteryx, Taralea and Pterodon. This tribe may form an outlying group to the Dalbergieae, although Polhill (l981b) considered the nearest affinity to be with Monopteryx in the Dussia group in the tribe Sophoreae (Polhill 1981a). In this paper, the wood anatomy of the three genera is described and com pared with that of Sophoreae (Gasson 1994, Fujii et al. 1994), Swartzieae (Cowan 1981; Gasson 1996; Gasson & Webley 1999), and most genera of Dalbergieae. This work is part of a systematic study of the wood of all available genera in the Papil ionoideae, and will be followed by a survey of Dalbergieae wood anatomy. All three genera of Dipterygeae have previously been described separately in wood identification manuals, but there are some discrepancies between anatomy and taxonomy that are highlighted and discussed in this paper. Downloaded from Brill.com10/09/2021 10:01:17PM via free access 442 IAWA Journal, Vol. 20 (4), 1999 MATERIALS AND METHODS The descriptions are based on sections of the following wood sampIes: Dipteryx Schreb.: D. alata: SJRw 47807, Bolivia; SJRw 50210, Bolivia - D. ferrea: MADw 12465, Brazil; (D. cf. ferrea) MADw 22287, Peru- - D. lacunifera: MADw 45211, Brazil - D. magnifica: MADw 46441, Brazil; SJRw 22606 - D. micrantha: SJRw 36652, Brazil - D. panamensis: SJRw 45644, Panama; SJRw 3015 - D. polyphylla: SJRw 33835, Brazil; SJRw 42406, Venezuela - D. punctata: SJRw 34090, Brazil; SJRw 43591, Guyana; MADw 31887, Venezuela - D. rosea (probably a syno nym of D. odorata): MADw 31888, Venezuela. Pterodon Vogel: P abruptus: B.B. Klitgaard 75, Brazil - P emarginatus (labelIed P pubescens): MADw 18849, Bo1ivia. Taralea AubI.: T casiquiarensis: SJRw 42161, Venezuela - T crassifolia (labelIed D. crassifolia): MADw 41847 (= Uw 27320), Guyana - T oppositifolia: SJRw 23633, Brazil; SJRw 36631, Brazil; SJRw 36655, Brazil- T rigida (labelIed D, rigida): MADw 45218, Brazil; SJRw 54508, Venezuela. The names given here have been checked by Gwilym Lewis to ensure that the speci mens have been referred to the correct genus and species. GENERAL WOOD DESCRIPTION Growth rings not seen. Wood diffuse-porous. Vessels solitary, in pairs, some short ra dial multiples and clusters, but in no predominant pattern. Vessel elements relatively short, with alternate, fine (but very fine in Taralea species except for T. casiquiarensis), vestured intervascular pits and simple perforation plates. Vessel-ray pitting similar to intervascular pitting in size and shape. Fibres thick- to very thick-walled, non-septate with slit-like, apparently simple pits. Axial parenchyma scanty paratracheal to vasi centric, aliform and confluent, storied except in most Taralea species. Rays uniseriate or biseriate, three cells wide in a few species, mainly homocellular, consisting mostly of procumbent cells, storied in Dipteryx, Pterodon and T. casiquiarensis, but not storied in the other Taralea species. Prismatic crystals in chambered axial parenchyma cells present in most species, possibly also in some fibres adjacent to axial parenchyma. GENERIC WOOD DESCRIPTIONS AND PREVIOUS LITERATURE Dipteryx - Fig. 1-17 About 10 species from Central and South America, nine examined here. There are many publications with wood anatomical descriptions, including Anonymous (1981 a, 1981b), Pfeiffer (1926), Record & Mell (1924), Lindeman et al. (1963), Sudo (1963), Reinders Gouwentak & Rijsdijk (1968), Huizzi (1974), Mainieri (1978), Dechamps (1980), Gaiotti de Peralta & Edlmann Abbate (1981), Detienne et al. (1982), Detienne & Jacquet (1983), Mainieri et al. (1983), Tomazello Filho et al. (1983), Mainieri & Chimelo (1989), Ilic (1991), and Nardi-Berti & Edlmann Abbate (1992). Downloaded from Brill.com10/09/2021 10:01:17PM via free access Gasson - Anatomy of the Dipterygeae (Leguminosae) 443 Vessels solitary, in pairs and radial multiples, occasionally up to 7 (Fig. 3), some with gummy or resinous deposits. Intervessel pitting fine (up to c.l 0 /illl), clearly vestured. Fibres mostly very thick-walled. Axial parenchyma scanty paratracheal to vasicentric, aliform and confluent, storied, strands 2-4-celled. Rays uniseriate, rarely biseriate and more rarely up to 3 cells wide (D.jerrea), usually up to c. 15 cells high, storied. Sometimes up to 25 or 35 cells high and axially fused in D. magnifica (MADw 46441). Pterodon - Fig. 18-21 Perhaps 6 species in Brazil and Bolivia, but critical reappraisal is needed (Polhill 1981 b). Only two species, P. abruptus and P. emarginatus were examined here. Sev eral publications contain wood descriptions, including Mainieri & Primo (1971), Detienne & Jacquet (1983), Mainieri et al. (1983), Mainieri & Chimelo (1989), and Nardi-Berti & Edlmann Abbate (1992). Vessels solitary and in frequent radial multiples, some clusters, some vessels with resinous or gummy deposits. Intervessel pitting fine (up to c. 10 /illl), clearly vestured. Fibres very thick-walled. Axial parenchyma mainly aliform and confluent, strands storied and 2-4-celled. Rays uniseriate, very occasionally biseriate, usually up to 12 cells high, but occasionally up to 22 cells high in P. emarginatus, storied. Prismatic crystals abundant in P. abruptus, not seen in P. emarginatus. Taralea - Fig. 22-33 About 5 species from tropical South America. Four species were examined here. Taralea oppositijolia is described and illustrated by Anonymous (1981a), as showing storied rays and axial parenchyma as seen in Dipteryx and Pterodon. Detienne et al. (1982) also describe T. oppositijolia (as Dipteryx oppositijolia), not mentioning storeying, although they recognise it in D. odorata and D. punctata. Detienne & J acquet (1983) illustrate D. oppositijolia, which is not storied, presumably from the same wood sample(s). Evidence for storeying in T. oppositijolia is therefore equivocal, although in view of the non-storied nature of all Taralea species examined here (ex cept for T. casiquiarensis, see below), it seems most likely that the T. oppositijolia in Anonymous (1981a) is a misidentified Dipteryx or Pterodon. Vessels solitary, in pairs and radial multiples up to at least 6 in T. oppositijolia. Some vessels with resinous or gummy deposits. Intervessel pitting very fine (up to c. 4/illl). Fibres very thick-walled, most with no lumen. Axial parenchyma scanty paratracheal, aliform to confluent. Most axial parenchyma cells in T. rigida are sclerified (Fig. 27). Axial parenchyma strands 2-4-celled, irregularly or not storied. Rays uniseriate and occasionally biseriate, often less than 20 cells high, but up to 38 cells high in T. oppo sitijolia, not storied. Some rays in T. oppositijolia have one row of upright marginal cells. Taralea casiquiarensis differs from the other Taralea species in having much larger intervessel pitting (up to 10.5 /illl) and storied axial parenchyma and rays. (text continued on page 452) Downloaded from Brill.com10/09/2021 10:01:17PM via free access 444 IAWA Journal, Val. 20 (4), 1999 Fig. 1-4. Dipteryx. All TS. - D. alata, SJRw 50210 (1), D. ferrea, MADw 12465 (2), D. lacunifera (3), D. polyphylla, SJRw 42406 (4). - Scale line is 200 f.UI1 für all. Downloaded from Brill.com10/09/2021 10:01:17PM via free access Gasson - Anatomy of the Dipterygeae (Leguminosae) 445 Fig. 5 - 8. Dipteryx. All TS. - D. magnifica, MADw 46441 (5), D. micrantha (6), D. panamen sis, SJRw 45644 (7), D. punctata, SJRw 34090 (8). - Scale line is 200 ~ for all. Downloaded from Brill.com10/09/2021 10:01:17PM via free access 446 IAWA Journal, Vol. 20 (4), 1999 Fig, 9-13. Dipteryx. -D. rosea, TS (9), D. rosea, TLS (10), D. magnifica, MADw 46441, TLS (11), D. magnifica, SJRw 22606, TLS (12), D. alata, SJRw 50210, TLS (13). - Scale line is 200 /l111 for all. Downloaded from Brill.com10/09/2021 10:01:17PM via free access Gasson - Anatomy of the Dipterygeae (Leguminosae) 447 Fig. 14-17. Dipteryx. - D. polyphylla, SJRw 42406, TS (14), D. cf.jerrea, MADw 22287, RLS (15), D. alata, SJRw 50210, TLS (16), D. magnifica, MADw 46441, TLS (17). - Scale line is 200 lJIIl for Fig. 14 & 15,50 lJIIl for Fig. 16 & 17. Downloaded from Brill.com10/09/2021 10:01:17PM via free access 448 IAWA Journal, Vol. 20 (4), 1999 Fig. 18-21. Pterodon emarginatus (= P. pubescens). - TS (18), TLS (19), RLS (20), TLS (21). - Scale line is 200 JllIl for Fig. 18-20, 50 JllIl for Fig. 21. Downloaded from Brill.com10/09/2021 10:01:17PM via free access Gasson - Anatomy of the Dipterygeae (Leguminosae) 449 Fig. 22-25. Taralea casiquiarensis. - TS (22), TLS (23), RLS (24), TLS (25). - Scale line is 200 llID für Fig. 22-24, 50 llID for Fig. 25. - This species is belng reassigned tü Dipteryx by Lewis & Gassün (in press). Downloaded from Brill.com10/09/2021 10:01:17PM via free access 450 IAWA Journal, Vol. 20 (4), 1999 Fig. 26-29. Taralea. T. crassifolia (26, 28, 29), T. rigida, MADw 45218 (27). - TS (26, 27), TLS (28, 29).